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Thelypteridaceae is a family of about 900 species of
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except ...
s in the order
Polypodiales The order Polypodiales encompasses the major lineages of polypod ferns, which comprise more than 80% of today's fern species. They are found in many parts of the world including tropical, semitropical and temperate areas. Description Polypodiale ...
. In the
Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group, or PPG, is an informal international group of systematic botanists who collaborate to establish a consensus on the classification of pteridophytes (lycophytes and ferns) that reflects knowledge about plant relatio ...
classification of 2016 (PPG I), it is placed in the suborder Aspleniineae. Alternatively, the family may be submerged in a very broadly defined family Aspleniaceae as the subfamily Thelypteridoideae. The ferns are
terrestrial Terrestrial refers to things related to land or the planet Earth. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
, with the exception of a few which are
lithophyte Lithophytes are plants that grow in or on rocks. They can be classified as either epilithic (or epipetric) or endolithic; epilithic lithophytes grow on the surfaces of rocks, while endolithic lithophytes grow in the crevices of rocks (and are als ...
s (grow on rocks). The bulk of the species are tropical, although there are a number of temperate species. These ferns typically have creeping rhizomes. The
frond A frond is a large, divided leaf. In both common usage and botanical nomenclature, the leaves of ferns are referred to as fronds and some botanists restrict the term to this group. Other botanists allow the term frond to also apply to the lar ...
s are simply
pinnate Pinnation (also called pennation) is the arrangement of feather-like or multi-divided features arising from both sides of a common axis. Pinnation occurs in biological morphology, in crystals, such as some forms of ice or metal crystals, an ...
to pinnate- pinnatifid. There is either no frond
dimorphism Dimorphism or dimorphic may refer to: Science * Dimorphic root systems, plant roots with two distinctive forms for two separate functions * Sexual dimorphism, a phenotypic difference between males and females of the same species * Nuclear dimorph ...
or only mild dimorphism, either open venation or very simple anastomosing. The sori are mostly reniform in shape and have indusia, except for the ''Phegopteris'' group.


Classification

During the early and mid 1900's all thelypterioid ferns were included in the genus '' Dryopteris'' because of the sorus shape. However, there are a great many differences between the groups, and these plants are now segregated in their own family. Genetic evidence shows that the family is clearly
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic ...
.


Phylogenetic relationships

The following cladogram for the suborder Aspleniineae (as eupolypods II), based on Lehtonen (2011), and Rothfels & al. (2012), shows a likely
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
relationship between the Thelypteridaceae and the other families of the clade.


Genera

The family can be divided into two major clades, which the
Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group, or PPG, is an informal international group of systematic botanists who collaborate to establish a consensus on the classification of pteridophytes (lycophytes and ferns) that reflects knowledge about plant relatio ...
classification of 2016 (PPG I) recognizes as two subfamilies, Phegopteridoideae and Thelypteroideae. Their division into genera has been described as "highly controversial and fluctuating". The family includes several complexes of species that are difficult to distinguish, and seem to represent a remarkable evolutionary radiation. Some researchers include the entire family Thelypteridaceae in the genus ''
Thelypteris ''Thelypteris'' (maiden ferns) is a genus of ferns in the subfamily Thelypteridoideae, family Thelypteridaceae, order Polypodiales. Two radically different circumscriptions of the genus are in use . In the Pteridophyte Phylogeny Group classific ...
''; others divide the family into as many as 30 genera. An intermediate position is to place the bulk in ''Thelypteris'' (which can then be divided into subgenera and sections corresponding to the genera of other authors) but to separate out ''
Phegopteris ''Phegopteris'' is a genus of ferns in the family Thelypteridaceae, subfamily Phegopteridoideae, in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). They are known collectively as the beech ferns. Species are native to Asia, N ...
'' and ''
Macrothelypteris ''Macrothelypteris'' is a genus of ferns in the family Thelypteridaceae, subfamily Phegopteridoideae, in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). The genus name means 'large ''Thelypteris. Species , the ''Checklist of ...
''. Another choice is to divide the family into a half a dozen or so genera. The Pteridophyte Phylogeny Group classification of 2016 (PPG I) accepts 30 genera: *Subfamily Phegopteridoideae Salino, A.R.Sm. & T.E.Almeid **''
Macrothelypteris ''Macrothelypteris'' is a genus of ferns in the family Thelypteridaceae, subfamily Phegopteridoideae, in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). The genus name means 'large ''Thelypteris. Species , the ''Checklist of ...
'' (H.Ito) Ching **''
Phegopteris ''Phegopteris'' is a genus of ferns in the family Thelypteridaceae, subfamily Phegopteridoideae, in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). They are known collectively as the beech ferns. Species are native to Asia, N ...
'' (C.Presl) Fée **''Pseudophegopteris'' Ching – here treated as a synonym of ''Phegopteris'' *Subfamily Thelypteridoideae C.F.Reed **''
Amauropelta ''Amauropelta'' is a genus of ferns in the family Thelypteridaceae, subfamily Thelypteridoideae, in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Other sources sink the genus into a very broadly defined genus ''Thelypteris''. ...
'' Kunze **'' Amblovenatum'' J.P.Roux **'' Ampelopteris'' Kunze **'' Chingia'' Holttum **''
Christella ''Christella'' is a genus of around 70-80 species of ferns in the subfamily Thelypteridoideae of the family Thelypteridaceae in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Other sources sink ''Christella'' into a very broadly ...
'' H.Lév. **''
Coryphopteris ''Coryphopteris'' is a genus of ferns in the family Thelypteridaceae, subfamily Thelypteridoideae, in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Species , the ''Checklist of Ferns and Lycophytes of the World'' accepted the ...
'' Holttum **''
Cyclogramma ''Cyclogramma'' is a genus of ferns in the family Thelypteridaceae, subfamily Thelypteridoideae, in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Other sources sink ''Cyclogramma'' into a very broadly defined genus ''Thelypter ...
'' Tagawa **''
Cyclosorus ''Cyclosorus'' is a genus of ferns in the family Thelypteridaceae, subfamily Thelypteridoideae, in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Other sources sink ''Cyclosorus'' into a very broadly defined genus ''Thelypter ...
'' Link **'' Glaphyropteridopsis'' Ching **'' Goniopteris'' C.Presl **'' Meniscium'' Schreb. **'' Menisorus'' Alston **'' Mesophlebion'' Holttum **'' Mesopteris'' Ching **'' Metathelypteris'' (H.Ito) Ching **'' Nannothelypteris'' Holttum **''
Oreopteris ''Oreopteris'' is a genus of ferns belonging to the family Thelypteridaceae. The species of this genus are found in Eurasia and Northern America. Species: * '' Oreopteris elwesii'' (Baker ex Hook. & Baker) Holttum * ''Oreopteris limbosperma ...
'' Holub **''
Parathelypteris ''Parathelypteris'' is a genus of ferns in the family Thelypteridaceae, subfamily Thelypteridoideae, in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Other sources sink ''Parathelypteris'' into a very broadly defined genus ''Th ...
'' (H.Ito) Ching **'' Plesioneuron'' (Holttum) Holttum **''
Pneumatopteris ''Pneumatopteris'' is a genus of about 80 species of terrestrial ferns in the family Thelypteridaceae. The range of the genus extends from tropical Africa, through Asia, Malesia and Australia to the Pacific islands Collectively call ...
'' Nakai **''
Pronephrium ''Pronephrium'' is a genus of ferns in the family Thelypteridaceae in the family Thelypteridaceae, subfamily Thelypteridoideae, in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Other sources sink ''Pronephrium'' into a very bro ...
'' C.Presl **'' Pseudocyclosorus'' Ching **'' Sphaerostephanos'' J.Sm. **''
Stegnogramma ''Stegnogramma'' is a genus of ferns in the subfamily Thelypteridoideae of the family Thelypteridaceae in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Other sources sink ''Stegnogramma'' into a very broadly defined genus ''The ...
'' Blume **'' Steiropteris'' (C.Chr.) Pic.Serm. **''
Thelypteris ''Thelypteris'' (maiden ferns) is a genus of ferns in the subfamily Thelypteridoideae, family Thelypteridaceae, order Polypodiales. Two radically different circumscriptions of the genus are in use . In the Pteridophyte Phylogeny Group classific ...
'' Schmidel **'' Trigonospora'' Holttum Some species of ''Pronephrium'' were split off into other genera in 2021 as a result of a
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
study of the family Thelypteridaceae: *''
Abacopteris ''Abacopteris'' is a small genus of ferns in the family Thelypteridaceae. Taxonomy ''Abacopteris'' was previously sunk into ''Pronephrium'' but was restored as a full genus as a result of a phylogenetic In biology, phylogenetics (; from Greek ...
'' Fée *''
Grypothrix ''Grypothrix'' is a small genus of ferns in the family Thelypteridaceae. Taxonomy ''Grypothrix'' was previously sunk into '' Pronephrium'' but was restored as a full genus as a result of a phylogenetic In biology, phylogenetics (; from Greek ...
'' (Holttum) S.E.Fawc. & A.R.Sm. *''
Menisciopsis ''Menisciopsis'' is a small genus of ferns in the family Thelypteridaceae. Taxonomy ''Menisciopsis'' was raised to the rank of genus from '' Pronephrium'' sect. ''Menisciopsis'' in 2021 as a result of a phylogenetic In biology, phylogenetics ( ...
'' (Holttum) S.E.Fawc. & A.R.Sm The extinct genus '' Holttumopteris'' from the Cenomanian aged
Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. ...
has been suggested to have affinities with the family, but several important diagnostic characters are not preserved.


References

{{Taxonbar, from=Q1496479 Fern families